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Dog Behaviour, Evolution, and Cognition

i
This page intentionally left blank
Dog Behaviour,
Evolution, and
Cognition

Ádám Miklósi
Department of Ethology, Eótvös Loránd University, Budapest

OXTORD
UNIVERSITY PRESS
OXPORD
UNIVERSITY PRESS
Great Clarendon Street, Oxford OX2 6DP
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Oxford is a registered trade mark of Oxford University Press
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Database right Oxford University Press (maker)
First published 2007
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
without the prior permission in writing of Oxford University Press,
or as expressly permitted by law, or under terms agreed with the appropriate
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Oxford University Press, at the address above
You must not circulate this book in any other binding or cover
and you must impose the same condition on any acquirer
British Library Cataloguing in Publication Data
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Library of Congress Cataloging in Publication Data
Data available
Typeset by Newgen Imaging Systems (P) Ltd., Chennai, India
Printed in Great Britain
on acid-free paper by
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ISBN 978-0-19-929585-2 (Hbk)
10 9 8 7 6 5 4 3 2 1
To my mother and father who have always believed that
I can do it, and to Zsuzsanka, Betty, and Gergo
who made doing it possible.
 Prologue: comparare necesse est*

In 1994, after some discussion, we decided to clear when Karin Grossman, a famous German child
our laboratories of the aquaria that had been in use psychologist, introduced us to Ainsworth's Strange
for many years in a research programme on the Situation Test, which is used to describe the pattern
ethology of learning in the paradise fish (Csanyi of attachment in children. Watching the videos on
1993). To be honest, the exact reason for this move how the children behaved when a stranger entered
at that time was not exactly clear to me, but I had the observation room or when their mother left,
no great regret for the research topic because we made us each realize independently that dogs
were the only laboratory studying learning pro- would behave in just the same way!
cesses associated with antipredator behaviour in It took us another two years to publish our first
this little East Asian labyrinth fish. study on the behavioural analysis of dog-human
However, the idea of approaching dog-human relationships based on the Strange Situation Test in
social interactions from an ethological perspective the Journal of Comparative Psychology, but from that
did not seem to be much of an improvement in that time on we had a quite clear idea of our research
respect, because literature on the subject was sim- programme, which was focused on looking for
ply non-existent. Thus József Topál, my colleague behavioural parallels between dogs and humans.
and friend, and I were a bit uncertain about the Actually, the idea of behavioural similarity
future when Professor Vilmos Csányi, the head of between humans and dogs was not novel at all.
the department at that time, began to argue enthu- Scott and Fuller (1965) devoted a considerable part
siastically that the study of dog behaviour in the of their work to human and dog parallels. For
human social context could be very important in example, in the first paragraph of their last chapter
understanding cognitive evolution, with many par- they write: These facts suggest a hypothesis: the
allels to human behaviour (Csányi 2000). We were genetic consequences of civilized living should
told hundreds of causal observations of dog-human be intensified in the dog, and therefore the dog
interaction (many people would call these anec- should give us some idea of the genetic future of
dotes), and it seemed that the task would be to pro- mankind....' In retrospect it is interesting that
vide an observational and experimental background although the achievements of this research group
to these ideas. Csányi pointed out that in order to be have always been recognized at the highest level,
successful in the human social world dogs had to these conclusions were neither debated nor praised
achieve some sort of social understanding, and very (or, more importantly, followed up in research).
likely this came about in course of their evolution. However, one point is important: although Scott
Accordingly, the social skills of dogs can be set in and Fuller realized the special social status of dogs
parallel with corresponding social skills in early in human groups in their behavioural work, they
humans. I do not know what exactly József thought emphasized parallels between the dog puppy and
about all this, but at least he owned a dog. the human child. In contrast, our aim was to pro-
After some thinking about what to do and how vide an evolutionary framework that hypothesizes
to do it, we saw some light at the end of the tunnel behavioural convergence between the two species.

*Comparison is essential; analogous to the Latin motto navigare necesse est, which can be translated as 'trade is essential'.

vi
 P R O L O G U E : COMPARARE NECESSE EST vii

Accordingly, we argued that evolutionary selective comparative perspective. This gave us the idea of
pressures for dogs might have moulded their socializing some wolves (and also some dog pups)
behaviour in such a way that it became compatible in order to obtain comparative data. This research
with human behaviour. not only opened our eyes to the very different
Since then, 12 years have passed and during that world of 'wild' canids but also taught us to be very
time many research groups have started to study cautious about coming to hasty conclusions about
dog behaviour. Although we have continued to behavioural differences between dog and wolf.
work according to our research programme, we Naturally, observations on these two species sug-
have realized that the field begs for integration. In gested many differences; however, the real trick
recent years many books on dogs have been pub- was to find the ways in which these differences
lished by researchers working in various fields, as could come to light under the conditions of a scien-
well as by experts with different backgrounds. The tific experiment. Later this comparative work was
goal of most of these books was to explain dog broadened to include cats and horses, but first of all
behaviour from an author's particular point of human children. We believe strongly that dog
view, often based on an assorted array of argu- behaviour can be understood only if it is studied in
ments where scientific facts were often treated at a comparative framework that takes into account
the same level as anecdotes, stories, or second-hand evolutionary and ecological factors and rests on a
information. In this book I want to break this solid methodological basis.
mould by presenting only what we know about Today, research inspired by ethology or behav-
dog behaviour and suggesting possible directions ioural ecology is characterized by a functional per-
for future research. The main aim is to provide a spective. Researchers focus their interest on those
common platform for scientific thinking for aspects of behaviour that contribute to the survival
researchers coming from the diverse fields of of the species. In the present case the focus is on a
archeozoology, anthrozoology, genetics, ethology, species, dogs, and on how collaboration among
psychology, and zoology. different scientific disciplines can lead to a more
The increased amount of contemporary research complete understanding of their evolution and pre-
has made it impossible to refer extensively to older sent state. For many years scientists have looked
work, much of which is, however, available in other with suspicion at dogs and denied them the status
textbooks. For similar reasons I have omitted to of 'real' animals. Thus the main goal of this book is
mention research that is not published in refereed to provide evidence that dogs can be studied just as
journals, or the many folk beliefs about dogs. In well as other animals (including humans) and even
addition, there is no attempt to 'bridge' gaps in our that they have the potential to become one of the
knowledge by 'facts that everyone knows', in the most well-researched species in the near future. In
absence of published evidence. Some readers may this regard dog ethology could play a role in pro-
see this as a serious fault which makes the presen- viding raw material for disciplines that are study-
tation of the topic uneven, but I have preferred to ing genetic and physiological aspects of behaviour,
use these opportunities to indicate directions in and also for those who are interested in applied
which research should be pursued. aspects such as dog training, problem behaviour,
Perhaps this is not the first book on dog ethology, dog-human interaction, or the use of dogs in thera-
but it has been written with the intent to place this peutic intervention.
species (once again) in the front line of ethology, I am very lucky to be a member of a wonderful
which is the science of studying animal (and research team with colleagues who have always
human) behaviour in nature. From the start we been supportive. I am grateful to Vilmos Csányi
believed that the whole project makes sense only if who gave us all the opportunity to embark on this
dogs are studied in their natural environment research programme. Over the years Jozsef Topál
where they share their life with humans in small became the best colleague and friend that one
or large groups. But we soon felt that such an could wish for in collaborative work, without
endeavour can only be insightful if it is put in a whom I would never have had the chance to get
viii P R O L O G U E : COMPARARE NECESSE EST

this project started. I owe a lot to Márta Gácsi who Gacsi (if not indicated otherwise). She and Eniko
has gently helped me in coming to understand the Kubinyi also made great efforts to help reading the
'world of dogs' over the years. I will never forget proof.
our first (and only) visit to Crufts. Antal Dóka, I would also like to thank to Richard Andrew,
who has been an indispensable colleague without Colin Allen, Laszlo Bartosiewitcz, Vilmos Csanyi,
whom the research group could not have func- Dorit Feddersen-Petersen, Simon Gabois, Marta
tioned so smoothly. Over the years we were lucky Gacsi, Borbala Gyori, Eniko Kubinyi, Daniel Mills,
to have Eniko Kubinyi, Zsófia Virányi, and Peter Eugenia Natali, Justine Philips, Peter Slater, Jozsef
Pongracz join our group, all of whom have made Topal, Judit Vas and Deborah Wells for reading
important contributions in particular fields of dog and commenting on single chapters or the whole
social behaviour and cognition. manuscript. Although these colleagues did every-
Over the years our research was supported by thing in their power to point out my weaknesses, I
the Eotvos Lorand University, the Hungarian shall take the responsibility for any mistakes left
Scientific Research Fund (OTKA), the Hungarian in the book.
Academy of Sciences, the European Union, the I am also grateful to Oxford University Press and
Ministry of Health, and the Dogs for Humans in particular to Ian Sherman for taking on this
Foundation. project without much hesitation, and also helping
Our research group owes much to those enthusi- to polish my raw Hungarian version of English.
astic dog owners and their dogs, who contributed Finally, a note to the critical reader. Please do not
by offering their time for our research. In addition hesitate to point out the weaknesses of this book.
we would like to express our thanks to Zoltan Not only to make the next version even better, but
Horkai and to the keen students (Bea Belenyi, also to urge others to provide facts in the form of
Eniko Kubinyi, Anita Kurys, Dorottya Ujfalussy, well-designed experiments that will separate sci-
Dorottya Ujvari, Zsofia Viranyi) who participated entific knowledge from beliefs and stories. If
in the Family Wolf Project and persisted in doing researchers and many others interested in dogs are
this job under difficult conditions. provoked to do better research then the book and I
I am very grateful to Antal Doka for drawing have achieved our goal.
and redrawing many figures and graphics for the
book. Being untalented at producing pictures, I am Budapest, 2 February 2007
thankful for the photos that were shot by Marta Adam Miklosi
 Contents

1 Dogs in historical perspective, and conceptual issues of

the study of their behaviour 1
1.1 Introduction 1
1.2 From behaviourism to cognitive ethology 2
1.2.1 Dog heroes visit the laboratory 3
1.2.2 Dogs in the comparative psychology laboratory 4
1.2.3 Naturalistic experiments 5
1.2.4 Time for comparisons 7
1.2.5 The cognitive revolution hits dogs 8
1.3 Tinbergen's legacy: four questions plus one 8
1.3.1 Description of behaviour 8
1.3.2 The first question: function 10
1.3.3 The second question: mechanism 10
1.3.4 The third question: development 10
1.3.5 The fourth question: evolution 11
1.4 Evolutionary considerations 11
1.5 What is it like to be a dog? 15
1.6 Lupomorphism or babymorphism? 16
1.7 Modelling of behaviour 17
1.7.1 Top down or bottom up 18
1.7.2 Canon of parsimony 18
1.7.3 Associanism and mentalism 19
1.7.4 Comparing content and operation 19
1.7.5 Comparing intelligence 21
1.7.6 Epigcncsis, socialization and cnculturation 22
1.8 An ethocognitive mental model for the dog 22
1.9 Conclusions for the future 24
Further reading 26

2 Methodological issues in the behavioural study of the dog 27

2.1 Introduction 27
2.2 Finding phenomena and collecting data 27
2.2.1 Qualitative description 28
2.2.2 Quantitative description 28
2.3 Making behavioural comparisons 30
2.3.1 Wolves and dogs 30
2.3.2 The comparison of breeds 33
2.3.3 Dogs and children 34
ix
x CONTENTS

2.4 Sampling and the problem of single cases (N = 1) 35

2.5 A procedural problem in naturalistic observations: the presence of humans 37
2.6 How to measure dog behaviour? 38
2.7 Asking questions 43
2.8 Conclusions for the future 45
Further reading 45

3 Dogs in anthropogenic environments: society and family 47

3.1 Introduction 47
3.2 Dogs in human society 47
3.3 Interactions between dogs and people in public 51
3.4 Dogs in the family 53
3.5 Dogs at work 56
3.6 Social roles of dogs in human groups 56
3.7 Social competition in dog-human groups and their consequences 57
3.7.1 Aggression and the human family 58
3.7.2 Studying the 'biting dog' phenomenon 58
3.7.3 Identifying risks 59
3.8 Outcast dogs: life in animal shelters 62
3.9 Conclusions for the future 65
Further reading 65

4 A comparative approach to Canis 67

4.1 Introduction 67
4.2 Putting things into perspective: an overview of Canis 67
4.2.1 Systematic relationships and geographic distribution 67
4.2.2 The evolution of Canis 68
4.2.3 The ecology and dynamics of group living in some canids 71
4.3 An overview of wolves 74
4.3.1 Geographic distribution and systematic relationships 75
4.3.2 Evolution of the wolf 76
4.3.3 Behavioural ecological aspects 79
4.3.4 Social relationships between and within wolf packs 81
4.3.5 A comparison: social organization in free-ranging dogs 86
4.4 Wolf and dog: similarities and differences 89
4.4.1 Morphological traits 90
4.4.2 Behavioural comparisons 90
4.5 Conclusions for the future 92
Further reading 93

5 Domestication 95
5.1 Introduction 95
5.2 Human perspective on dog domestication 95
5.3 Archaeology faces phylogenetics 101
5.3.1 The archaeologists' story: looking at archaeological evidence 101
5.3.2 The geneticists' story: evolutionary genetic evidence 109
 CONTENTS xi

5.4 Some concepts of evolutionary population biology 117

5.4.1 The question of founder population(s) 117
5.4.2 On the nature of selection 118
5.4.3 Changes in reproductive strategy and effects on generation times 119
5.5 Emergence of phenotypic novelty 119
5.5.1 Mutation 120
5.5.2 Hybridization 123
5.5.3 Directional trait selection 124
5.5.4 Selection for plastic phenotypes 125
5.5.5 Heterochrony 126
5.5.6 The 'mysterious laws' of correlation 129
5.6 A case study of domestication: the fox experiment 131
5.6.1 The founding foxes and behavioural selection 131
5.6.2 Changes in early development 133
5.6.3 Changes in the reproductive cycle 134
5.6.4 Have we got domesticated foxes? 135
5.7 Conclusions for the future 136
Further reading 136

6 The perceptual world of the dog 137

6.1 Introduction 137
6.2 Comparative perspectives 137
6.2.1 Cognitive aspects of perception 138
6.2.2 Experimental approach to study perceptual abilities 139
6.3 Vision 139
6.3.1 Physical processing 139
6.3.2 Neural processing and visual ability 141
6.3.3 Perception of complex visual images 142
6.4 Hearing 142
6.4.1 Physical processing 142
6.4.2 Neural processing and hearing ability 142
6.4.3 Perception of complex sound forms 143
6.5 Olfaction 144
6.5.1 Physical processing 144
6.5.2 Neural processing and olfactory ability 144
6.5.3 Categorization and matching in working situation 147
6.5.4 Perception of natural substances and conspecific odours 149
6.6 Conclusions for the future 150
Further reading 150

7 Physical-ecological cognition 151

7.1 Introduction 151
7.2 Orientation in space 151
7.2.1 Path following 152
7.2.2 Beacons 152
7.2.3 Landmarks 152
7.2.4 Egocentric orientation 154
xii CONTENTS

7.3 Spatial problem solving 155

7.4 Knowledge about objects 156
7.5 Memory for hidden objects 158
7.6 Folk physics in dogs? 160
7.6.1 Means-end connections 161
7.6.2 'Gravity' 162
7.7 Conclusions for the future 162
Further reading 163

8 Social cognition 165

8.1 Introduction 165
8.2 The affiliative aspects of social relationships 166
8.3 The agonistic aspects of social relationships 170
8.3.1 Classification of aggression in dogs 172
8.3.2 Is there an ethological description of aggressive behaviour in dogs? 172
8.3.3 Decreased aggression in dogs? 173
8.3.4 Organization of aggressive behaviour and the role of learning 173
8.3.5 Reaction to human agonistic signals 174
8.4 Communication in a mixed-species group 177
8.4.1 Visual communication 178
8.4.2 Acoustic communication 185
8.5 Play 189
8.6 Social learning in dogs 191
8.7 Social influence 193
8.8 Cooperation 196
8.9 Social competence 197
8.10 Conclusions for the future 200
Further reading 200

9 Development of behaviour 201

9.1 Introduction 201
9.2 What are developmental 'periods'? 201
9.3 Rethinking developmental periods in dogs 205
9.3.1 Neonatal period 206
9.3.2 Transition period 206
9.3.3 Socialization period 207
9.3.4 Juvenile period 208
9.4 Sensitive periods in development 209
9.5 Attraction and attachment 214
9.6 Early experience and its influence on behaviour 216
9.7 Prediction of behaviour:'Puppy testing' 217
9.8 Conclusions for the future 219
Further reading 219

10 Temperament and personality 221

10.1 Introduction 221
 CONTENTS xiii

10.2 Descriptive approach to personality 223

10.2.1 'Knowing', observing, or testing 223
10.2.2 Describing behaviour: assessment and coding 224
10.2.3 The construction of personality 225
10.3 Functional approach to personality 226
10.4 Mechanistic approach 230
10.4.1 Insights from genetics 230
10.4.2 Physiological correlates of personality traits 231
10.5 Conclusions for the future 234
Further reading 235

11 Afterword: Heading towards 21st-century science 237

11.1 Comparare necesse est! 237
11.2 Natural model 237
11.3 Evolving dogs 237
11.4 Behavioural modelling 239
11.5 Ethical implications and researchers' mission 240
11.6 Dog genome and bioinformatics 241
11.7 Taws in hands' 241

References 243
Index 267
This page intentionally left blank
 CHAPTER 1

Dogs in historical perspective,

and conceptual issues of the
study of their behaviour

1.1 Introduction The study of dogs did not fit well with the
increasing influence of behavioural ecology, which
This book is about the biological study of dog behav- was partially initiated by the call for a more func-
iour, based on the programme summarized so clearly tional approach to behaviour by Tinbergen (1963).
by Tinbergen in 1963. He, Lorenz and others have Obviously, dogs are not the best candidates for
always pointed out that the main contribution of eth- studying survival in nature, mainly because most
ology is the biological analysis of animal behaviour present-day dogs live with humans and have access
based on observations in nature. Unfortunately, to vets, and we do our best to save our companions
however, only a handful of mainstream ethologists from the challenges of nature. In this sense dogs
have applied these concepts to dog behaviour. In can be regarded as being special (but not necessar-
contrast to sticklebacks, honeybees or chimpanzees, ily 'artificial').
not to mention a few tens of other species, dogs More surprisingly, interest in the study of dogs
received relatively little attention from ethologists or did not emerge with the cognitive revolution in
comparative psychologists. It seems that these crea- ethology. Griffin (1984), one of the initiators of this
tures ('man's best friends') have somehow become movement, seems to have carefully avoided refer-
outcasts from mainstream science, for reasons that ence to dogs in most of his works on this subject.
are not obviously clear but which may be guessed. We are introduced to miraculous behaviour of ants,
Dogs are often referred to as 'artificial animals', starlings or dolphins, which we look at with admir-
probably because their history of being 'domesti- ation, but similar behaviour in dogs is often regarded
cated'. Here the image is that of a 'savage' stealing as suspicious. To some extent this attitude is under-
a wolf cub from its mother (e.g. Lorenz 1954), which standable, as early workers were often tricked by
then 'became' dog after many years and generations so called 'dog artists' who showed remarkable
in the hands of humans. Today most researchers skills for 'talking' or 'counting' (e.g. Pfungst 1912,
disagree with this simplistic view of dog domesti- Grzimek 1940-41). (Figure 1.1) After it was found out
cation (e.g. Herre and Rohrs 1990), and it is much that such apparently clever behaviour could be
less clear on what grounds the evolution of such explained by the dog responding to minute bodily
'real' and 'artificial' animals can be differentiated. cues produced either consciously or unconsciously
The kind of goal-directed selective breeding by the owner or trainer (the Clever Hans effect, see
implied by the category of 'artificial animal' prob- Pfungst 1907 and Chapter 2.5, p. 37), dogs were ban-
ably started much later than has been assumed. ished from laboratories for being unreliable subjects.
Logically, an 'artificial animal' cannot have a nat- However, it seems that dogs are showing signs of
ural environment, so in order to allow the dog into making a real comeback. Ethologists, comparative
the club of 'real' animals we would have to find a psychologists, and many others are now working
natural environment for it (Chapter 3, p. 42). hard to find a place for dogs in the biological study

1
 HISTORICAL PERSPECTIVE AND CONCEPTUAL ISSUES

Figure 1.1 (a) Stuppke, a counting dog artist, was observed by Bernhard Grzimek, a German zoologist. Stuppke barked the number shown
to him. The remarkable talent of the dog was based on recognition of a 'start' and a 'stop' signal given by his master, Mr Pilz. (b) No wonder
that Stuppke could also read numbers with his eyes covered (photos taken from Grzimek 1940-41). (c) Oskar Pfungst (1912) reported on Don,
the talking dog (photo from Candland 1993, Oxford University Press).

of behaviour. This is difficult, but the steep increase

in research papers over the last 10 years already
shows the fruit of this work. Thus there is every
chance that dog ethology will revive.

1.2 From behaviourism to

cognitive ethology
Early researchers, including Darwin (1872),
regarded the dog as a special animal that is com-
parable to humans. Many people shared this
anthropomorphic attitude and it is not surprising
that dogs ended up at the top of the ladder repre-
senting intelligence and emotional behaviour in
animals (Romanes 1882a, b) (Figure 1.2). It did
not take long for the situation to change, and
dogs could not avoid their fate when under the
increasing influence of behaviourism they were
then treated as a sort of stimulus-response
automaton. The interest in wolves and social
behaviour in general has helped dogs regain a
foothold in the behavioural sciences, and this has
The Dog at the Convent Door. led to an ethologically oriented understanding of
dog behaviour. The history of the study of dogs
Figure 1.2 The 'cultural transmission' of dog anecdotes. Menault reflects the changes in our views of animals, and
(1869) reports the story of a dog that, after observing beggars
although much time has passed and a lot of
ringing the bell at the door of the convent and receiving some soup,
went to the door and pulled the string. The ability to learn by
knowledge has been gained, the basic questions
observation of humans has only recently been demonstrated of present-day research are more or less the same
experimentally (Chapter 8, e.g. Kubinyi eta/. 2001 36; Box 8.6). as they were 100 years ago.
 1.2 FROM B E H A V I O U R I S M TO C O G N I T I V E E T H O L O G Y 3

1.2.1 Dog heroes visit the laboratory dogs were much inferior in escaping. From his
descriptions it seems that, in comparison with the
Dogs have long been the favourite heroes of
cats, dogs were less inclined to get out, and they
animal stories. Sharing our daily life with these
were also very cautious in interacting with the
animals has offered endless opportunities to
latch, which probably indicates a different social
observe or witness the varieties of dog-human
relation between people and these dogs. Thus it is
interactions. One famous collector of such stories
less surprising that in the textbooks the fame of
was George Romanes (1982a). His descriptions of
representing Thorndike's concept of trial-and-error
dogs provided evidence for often very intelligent
learning was left to the cats. From further experi-
behaviour which prompted him to argue that
ments Thorndike did not find support for the
such performances should be explained by
long-held view that dogs learn by imitation (see
human-like thinking mechanisms (Candland
Chapter 8.6, p. 191) because animals did not escape
1993).
any earlier from the box if they were shown how to
Interestingly, Lloyd Morgan (1903), who was a
open the lock.
strong critic of the methods used by Romanes, did
In 1904 Pavlov received the Nobel Prize for
not refrain from telling such stories when he
Medicine for the physiological study of the digest-
wanted to illustrate a particular behavioural phe-
ive system, for which dogs had served as subjects.
nomenon. At one point he describes how his fox
By this time he had noted that not just the presence
terrier Tony grappled with the problem of how to
of food in the mouth but also other external stimuli
carry a stick with unequal weights at its ends. After
(the sound of the food put in the bowl or the
describing the dog's behaviour Morgan concludes
approaching experimenter providing the food)
that he has seen little evidence for assuming that
have the potential to elicit salivation. For many
the dog 'understood the problem'. Instead, during
years after that dogs remained one of the most pre-
repeated attempts to carry the stick the dog learned
ferred subjects in the research that led to the devel-
the solution by trial and error. Thus 'intelligent'
opment of the conditioned reflex principle (Pavlov
behaviour on the dog's part could often be based
1927), which was extended by Pavlov's pupils.
on relative simple learning processes. For Morgan,
Pavlov was not only a good experimenter, however,
stories provided opportunities for formulating
but also a good observer. Thus he noted early on
hypotheses and did not serve as explanations for
that there are marked individual differences among
mental abilities. Nevertheless he did not deny that
the dogs, which could be also observed in their
dogs could have a mental representation for an
response to the training (Teplov 1964). Dogs were
object, such as a bone.
categorized as belonging to one of the classic tem-
Thorndike (1911) was among the first to develop
perament types described by Hippocrates (san-
a method to objectively measure learning in ani-
guine, choleric, phlegmatic, melancholic) (see also
mals. He put hungry cats and dogs into a box which
Box 10.1). Even at that time Pavlov pointed out that
could be opened from inside by manipulating a
observed behavioural traits are the outcome of
simple latch. Observing the animals repeatedly in
complex processes having both genetic and envir-
this situation, he found that it took them less and
onmental components, and he was probably the
less time to get out. In agreement with Morgan, he
first to suggest separating these two effects by rais-
also thought that the final 'intelligent' behavioural
ing dogs in different environments before subject-
solution was the result of a step-by-step process of
ing them to training. The generality of Pavlov's
'trial and error' learning. Thus the systematic
work on the conditioning reflexes provided the
observations of both Morgan and Thorndike
basis for comparative work on dogs and humans.
seemed to contradict the conclusions of Romanes,
Based on this experimental approach, dogs can be
who argued that, for example, cats and dogs have
regarded as the first animal models of human per-
someideaaboutthepropertiesof locks.Interestingly,
sonality (Chapter 10, p. 221). This makes it less sur-
Thorndike noted a difference between dogs and
prising that in contrast to some other laboratories
cats, because, despite being starved for some time,
Pavlov's researchers respected the individuality of
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